HORT640 - Metabolic Plant Physiology
Proline, ornithine and arginine metabolism
Pathway of proline synthesis in bacteria
In bacteria such as Escherichia coli, Pseudomonas aeruginosa and Salmonella typhimurium, proline synthesis from glutamate is catalyzed by three enzymes: gamma-glutamyl kinase (GK), glutamyl phosphate reductase (GPR) (glutamate-5-semialdehyde dehydrogenase), and pyrroline-5-carboxylate reductase (P5CR), encoded by genes proB, proA and proC, respectively (Baich, 1969; 1971; Gamper and Moses, 1974; Hayzer and Moses, 1978ab; Krishna and Leisinger, 1979; Krishna et al, 1979; Adams and Frank, 1980; Hayzer and Leisinger, 1980; 1981; 1982; Deutch et al, 1982; 1984; Smith et al, 1984):
Glutamyl kinase (glutamate 5-kinase) (GK) [EC 2.7.2.11]
Glutamate + ATP ---> glutamyl 5-phosphate + ADP
Glutamyl phosphate reductase (glutamic-5-semialdehyde dehydrogenase) (GPR) [EC 1.2.1.41]
glutamyl 5-phosphate + NADPH <---> glutamic-5-semialdehyde + NADP+ + Pi
Spontaneous
glutamic-5-semialdehyde <---> pyrroline-5-carboxylate +H2O
Pyrroline-5-carboxylate reductase (P5CR) [EC 1.5.1.2]
pyrroline-5-carboxylate + NADPH <---> proline + NADP+
The yeast proline biosynthesis pathway appears to be similar to the bacterial pathway (Li and Brandiss, 1992). The GK of Escherichia coli is inhibited by proline (Baich, 1969; Rushlow et al, 1984; Smith, 1985). Proline over-producing mutants have a GK with markedly altered feedback inhibition (Csonka, 1981; Rushlow et al, 1984; Smith, 1985).
References
Adams E, Frank L 1980 Metabolism of proline and the hydroxyprolines. Annu. Rev. Biochem. 49: 1005-1061.
Baich A 1969 Proline synthesis in Escherichia coli. A proline-inhibitable glutamic acid kinase. Biochim. Biophys. Acta 192: 462-467.
Baich A 1971 The biosynthesis of proline in Escherichia coli. Phosphate-dependent glutamate gamma-semialdehyde dehydrogenase (NADP), the second enzyme in the pathway. Biochim. et Biophys. Acta 244: 129-134.
Csonka LN 1981 Proline over-production results in enhanced osmotolerance in Salmonella typhimurium. Mol. Gen. Genet. 182: 82-86.
Deutch AH, Rushlow KE, Smith CJ, Kretschmer PJ 1982 Escherichia coli delta1-pyrroline-5-carboxylate reductase: gene sequence, protein overproduction and purification. Nucleic Acids Res. 10: 7701-7714.
Deutch AH, Smith CJ, Rushlow KE 1984 Analysis of the Escherichia coli proBA locus by DNA and protein sequencing. Nucleic Acids Res. 12: 6337-6355.
Gamper H, Moses V 1974 Enzyme organization in the proline biosynthetic pathway of Escherichia coli. Biochim. Biophys. Acta 354: 75-87.
Hayzer DJ, Leisinger T 1980 The gene-enzyme relationships of proline biosynthesis in Escherichia coli. J. Gen. Microbiol. 118: 287-293.
Hayzer DJ, Leisinger T 1981 Proline biosynthesis in Escherichia coli. Stoichiometry and end-product identification of the reaction catalyzed by glutamate semialdehyde dehydrogenase. Biochem. J. 197: 269-274.
Hayzer DJ, Leisinger T 1982 Proline biosynthesis in Escherichia coli. Purification and characterization of glutamate-semialdehyde dehydrogenase. Eur. J. Biochem. 121: 561-565.
Hayzer DJ, Moses V 1978a Proline biosynthesis by cell-free extracts of Escherichia coli and potential errors arising from the use of a bioradiological assay procedure. Biochem. J. 173: 207-217.
Hayzer DJ, Moses V 1978b The enzymes of proline biosynthesis in Escherichia coli. Their molecular weights and the problem of enzyme aggregation. Biochem. J. 173: 219-228.
Krishna RV, Beilstein P, Leisinger T 1979 Biosynthesis of proline in Pseudomonas aeruginosa. Properties of gamma-glutamyl phosphate reductase and delta1-pyrroline-5-carboxylate reductase. Biochem. J. 181: 223-230.
Krishna RV, Leisinger T 1979 Biosynthesis of proline in Pseudomonas aeruginosa. Partial purification and characterization of gamma-glutamyl kinase. Biochem. J. 181: 215-222.
Rushlow KE, Deutch AH, Smith CJ 1984 Identification of a mutation that relieves gamma-glutamyl kinase from allosteric feedback inhibition by proline. Gene 39: 109-112.
Smith LT 1985 Characterization of a gamma-glutamyl kinase from Escherichia coli that confers proline overproduction and osmotic tolerance. J. Bacteriol. 164: 1088-1093.
Smith CJ, Deutch AH, Rushlow KE 1984 Purification and characteristics of a gamma-glutamyl kinase involved in Escherichia coli proline biosynthesis. J. Bacteriol. 157: 545-551.
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